Intraspecific Phenotypic Variation in Nearly Threatened Mottled Nandus
Total Page:16
File Type:pdf, Size:1020Kb
Sains Malaysiana 49(11)(2020): 2609-2623 http://dx.doi.org/10.17576/jsm-2020-4911-01 Intraspecific Phenotypic Variation in Nearly Threatened Mottled Nandus,Nandus nandus (Hamilton, 1822) (Variasi Fenotip Intrakhusus Patung Belang Nandus yang Hampir Terancam, Nandus nandus (Hamilton, 1822)) MD. SAROWER-E-MAHFUJ, MD. ABDUS SAMAD, FEE FAYSAL AHMED, MD. ABDUL ALIM, YOSNI BAKAR & SIMON KUMAR DAS* ABSTRACT Understanding intraspecific phenotypic plasticity is a prerequisite of stock identification, evolutionary studies, sustainable utilization, and fishery conservation. In this study, intraspecific phenotypic plasticity was assessed in terms of the external features (i.e. meristic, morphometric, and truss-based morphometrics) of the wild Nandus populations from four freshwater sources in Southwestern Bangladesh. Fish samples were collected from Arial Kha River, Madaripur (AKRM, n=26); Nabaganga River, Jhenaidah (NRJ, n=22); Bohnni Baor, Gopalganj (BBG, n=26); and Dhakuria Beel, Jashore (DBJ, n=22). Meristic, morphometric, and truss network data were subjected to one-way ANOVA followed by post hoc (Tukey-HSD) test. The meristic counts of all the samples demonstrated significant differences only in one of the six characters. By contrast, significant differences were observed in 8 morphometric characters and 31 truss network data from 16 morphometric characters and 35 truss network data, respectively. Principal component (PCA) and canonical variate analyses (CVA) were also performed on morphometric and truss-based network data. Meristic and morphometric results from PCA and CVA showed that populations were completely intermingled, forming a compact cluster within intrapopulation levels, while truss morphometric characters formed a separate cluster. Three dendrograms independently based on phenotypic relationships among the individuals of the four populations also confirmed the absence of phenotypic differentiation among the population due to clustering of different groups. The baseline information resulting from the current study would be useful for genetic studies and further in situ conservation of Nandus populations in Bangladesh. Keywords: Canonical variate analysis; freshwater; morphometric; meristic; nandus; principle component analysis; Truss morphometry ABSTRAK Memahami keplastikan fenotip intrakhusus adalah satu pra-syarat untuk mengenal pasti stok, kajian evolusi dan pemanfaatan lestari dan pemuliharaan dalam perikanan. Dalam kajian ini kepelbagaian fenotip intrakhusus dinilai dari segi ciri luaran (iaitu meristik, morfometri dan morfometri dasarkan truss) daripada populasi liar ikan nandus dari empat sumber air tawar di selatan-barat Bangladesh. Sampel ikan dikumpulkan dari Arial Kha River, Madaripur (AKRM), (n = 26); Sungai Nabaganga, Jhenaidah (NRJ), (n = 22); Bohnni Baor, Gopalganj (BBJ), (n = 26); dan Dhakuria Beel, Jashore (DBJ), (n = 22). Data meristik, morfometri dan rangkaian truss dianalisis menggunakan varians satu arah (ANOVA) diikuti dengan ujian Post-hoc (Tukey-HSD). Perhitungan meristik untuk kesemua sampel menunjukkan perbezaan yang signifikan hanya dalam satu ciri daripada enam ciri manakala perbezaan yang signifikan diperhatikan dalam 8 ciri morfometrik dan 31 rangkaian data truss masing-masing daripada 16 ciri morfometrik dan 35 rangkaian data truss. Di samping itu, analisis komponen utama (PCA) dan analisis fungsi diskriminasi (CVA) dilakukan dengan menggunakan morfometrik dan data rangkaian berasaskan truss. Hasil daripada PCA dan CVA menunjukkan populasi terpisah sepenuhnya serta membentuk kelompok yang padat dalam tahap intra- INTRODUCTION populasi. Tiga dendrogram secara bebas berdasarkan hubungan fenotip antara individu daripada empat populasi River dibina. Populasi NRJ, BBG dan DBJ membentuk populasi kumpulan masing-masing berdasarkan meristik, morfometrik dan truss morfometrik. Maklumat asas yang dihasilkan daripada kajian semasa adalah mudah untuk kajian genetik dan pemuliharaan populasi Nandus secara in situ di Bangladesh. Kata kunci: Air tawar; analisis fungsi diskriminasi; meristik; morfometrik; morfometri Truss; nandus 2610 INTRODUCTION Nandus is a freshwater fish commonly known as mud perch or mottled nandus and considered a small Phenotypic plasticity is the ability of an organism to adjust indigenous species in Bangladesh (Ross et al. 2003). This its body maintenance in response to genetic-environmental fish species is widely distributed in fresh and brackish interactions. Sometimes, phenotypic plasticity, phenotypic waters, including ditches, ponds, beels (saucer-shaped responsiveness, flexibility, and condition sensitivity are perennial water bodies), and inundated fields throughout entirely synonymous in evolutionary biology (West- South Asian countries (Ahmed 2008; Rahman 2005). Eberhard 1989). The plethora of outcomes, such as changes Nandus is a carnivorous organism that entirely feeds on in body shape and size, allometry, feeding habits, sexual larvae and insects, crustaceans, filamentous algae, and dimorphism, and behavioral and physiological states, small fishes (Agarwal & Sharma 1966). Although this can be collectively or solely achieved from phenotypic species is considered a bony fish that survives at a low plasticity after a certain period of time (Langerhans oxygen level, it can camouflage when any prey, small fish, 2008). Thus, similar to other organisms with this and even a predator is present in a water body (Mustafa property, fishes are not an exception. Fishes also exhibit et al. 1980). This fish also plays a substantial role in the an outstanding extent of variation in their external body overall nutrition for poor-rural-living and low-income- shape morphologies, such as meristic and morphometric generating communities in Bangladesh (Das & Zamal characters, at a species level (Oufiero & Whitlow 2016). 2000). According to IUCN-Bangladesh (Chowdhury Consequently, morphometrics can be defined as an array of 2015), this species is categorized as nearly threatened quantitative analyses, such as biological outline, or shape because of habitat destruction, overexploitation, disparity among organisms with respect to environmental anthropogenic activities, and climate change (Rahman factors (Webster & Sheets 2010). Moreover, studies on 2005). As such, morphometric and meristic studies should the morphogenesis of fishes plays a fundamental role in be conducted to detect intraspecific phenotypic plasticity evolutionary analysis and proper management (Başusta et and ensure sustainability in the future. al. 2014; Kalhoro et al. 2015). At present, no adequate information regarding the Information related to the stock structure analysis of intraspecific phenotypic variation in N. nandus in the a species or a population is a prerequisite of the expansion freshwaters of Bangladesh is available. Therefore, this of proper biodiversity management and conservation study aimed to investigate the intraspecific phenotypic (Turan et al. 2005). Morphological dissimilarities are variations in N. nandus based on meristic, morphometric, observable characteristics in a fish or a fish population and truss network system. and caused by genetic factors, genetic-environmental interactions, and abiotic and biotic influences (Crispo MATERIALS AND METHODS 2008; Silva et al. 2013). Generally, in early developmental stages, fishes express their phenotypic plasticity in two FISH SAMPLING ways, that is, isometric size variation due to growth and allometric shape variation caused by developmental A total of 100 individuals of Nandus sp. were collected alteration (Cadrin 2000). Freshwater fishes exhibit a high from four different freshwater sources in Bangladesh degree of body shape variation because of physiological from September 2017 to November 2017: Arial Kha and environmental conditions, resulting in genetic River, Madaripur (AKRM); Nabaganga River, Jhenaidah variation and phenotypic plasticity (Eklöv & Svanbäck (NRJ); Bohnni Baor, Gopalganj (BBG); and Dhakuria 2005). Numerous techniques, such as morphometrics Beel, Jashore (DBJ) (Figure 1 & Table 1). The samples and meristics, traditional tags, otolith microchemistry, were placed in an ice box and immediately brought into and electronic tags, have been extensively used for stock the Laboratory of Fish Biology and Aquaculture, Jashore identification. Morphometric traits are one of the most University of Science and Technology, Bangladesh. The used and cost-effective methods to detect intraspecific minimum and maximum total lengths (TL) of the fish phenotypic variation in species (Mir et al. 2013). Naturally, specimens were 6.94 and 12.89 cm, respectively. fishes undergo ontogeny in an allometric pattern from the beginning of their life cycle (Hood & Heins 2000; Svanbäck COUNTING OF MERISTIC CHARACTERS & Eklöv 2002). To reinforce the inherent limitation of In six meristic characters, the numbers of dorsal spiny conventional morphometric approaches, the truss-networks fin rays (DSFR), dorsal soft fin rays (SFR), caudal fin rays formed by two or more interconnecting distances across- (CFR), anal fin rays (AFR), pelvic fin rays (PevFR), and body that ultimately produced chronological sequence pectoral fin rays(P ecFR) were counted in each sample by of associated polygons has been progressively utilized using magnifying glasses and needles. (Strauss & Bookstein 1982). 2611 FIGURE 1. Map of Bangladesh showing collection sites of N. nandus from four freshwater sources TABLE 1. Sampling details of N. nandus from four freshwater sources in Bangladesh Serial